Cryptogamie,Mycologie, 2008, 29 (2): 157-177 © 2008 Adac. Tous droits réservés

Records of French coprophilous fungi

Michael J. RICHARDSON

165 Braid Road,Edinburgh EH10 6JE U.K. [email protected] Telephone: 0131 447 8165

Résumé – Des détails sont donnés sur 169 espèces de mycètes coprophiles se développant sur 86 échantillons de fumier collectés en France (67 de la métropole, 5 de Corse, 5 de et 9 des îles Kerguelen) après incubation en chambres humides. ascomycètes / basidiomycetes / biogéographie / diversité / écologie / fimicoles

Abstract – Details are given of 169 species of coprophilous fungi developing on 86 samples of dung from France (67 from mainland France, 5 from , 5 from Guadeloupe and 9 from the Kerguelen Islands) and incubated in moist chambers. ascomycetes / basidiomycetes / biogeography / diversity / ecology / fimicoles

INTRODUCTION

During various visits to France from 1997-2007 samples of herbivore dung were collected and, on return to the UK, incubated in a damp chamber. Additionally, through the kind help of Jean-Louis Chapuis (Muséum national de Histoire naturelle, ), some samples were obtained from the Kerguelen Islands in the southern Indian Ocean between Australia and South Africa. The coprophilous fungi that developed were recorded.

MATERIAL AND METHODS

Details of the samples collected and incubated to provide records are given in Table 1, and their distribution is shown in Fig. 1. Most were dry when collected, and were placed in paper envelopes. Those that were not were air dried and then packeted. Most samples, other than those from the Kerguelen Islands, were rehydrated and incubated soon after collection on moist paper towelling in plastic boxes with lightly fitting transparent lids, under ambient light and at room temperature (ca 15-18°C). Care was taken to ensure that cultures were not too wet. Samples were generally of similar size, with incubation chambers 10 × 7 cm, which would accommodate approx. 2-4 g D.W. (= 15 sheep/goat/deer - 20 rabbit pellets), or 13 × 8 cm for horse or cattle (approx. 10 - 20 g D.W.). Samples were 158 M.J. Richardson

Fig. 1. Map showing the départements from which samples were collected, and the number from each. The two samples between and the Pyrénées-Orientales were collected on the boundary between the two départements. In addition, 5 samples were collected from Guadeloupe and 9 from the Kerguelen Islands. examined frequently at intervals of a few days, with a × 7 - 45 magnification stereomicroscope. Fruiting bodies were removed and mounted in water for examination and identification at higher magnification. Samples were mostly incubated for up to 22 wk (exceptionally for one sample for over a year), with observations continuing whilst new fungi were being observed. Sample locality coordinates (latitude and longitude) for most were determined with a Magellan GPS 4000 XL or eXplorist 100 GPS unit; those for Corsica and the Kerguelen Islands were obtained by using Google Earth and the locality details provided by the collectors. Selected material has been placed in the Herbarium of the Royal Botanic Garden,Edinburgh (E). In considering diversity, a cumulative species French coprophilous fungi 159

Table 1. Details of French dung samples and collection localities

Sample Elevation Latitude Longitude Locality Département Date Substrate no.* (m a.s.l.) (°) (°)

106/97 le Caroux Herault 1000 N43.59 E2.98 2.10.97 deer

107/97 le Caroux Herault 1000 N43.60 E2.98 2.10.97 hare

108/97 la Tour Herault 1050 N43.60 E3.00 2.10.97 deer

109/97 la Tour Herault 1050 N43.60 E3.00 2.10.97 hare

110/97 le Caylar Herault750 N43.88 E3.32 7.10.97sheep

111/97 800 N43.91 E3.32 7.10.97 hare

112/97 la Couvertoirade Aveyron 780 N43.90 E3.30 7.10.97sheep

113/97 la Couvertoirade Aveyron 780 N43.90 E3.30 7.10.97rabbit

114/97 le Caylar Herault725 N43.87 E3.30 7.10.97 cattle

115/97 Colombieres Herault740 N43.60 E3.018.10.97 deer

116/97 St Gervais Herault350 N43.66 E3.06 10.10.97 deer

117/97 St Gervais Herault350 N43.66 E3.06 10.10.97rabbit

18/98 Montgrand Tarn1240 N43.69 E2.72 20.4.98 hare

19/98 Roc de Montalet Tarn1240 N43.68 E2.74 20.4.98 cattle

20/98 Roc de Montalet Tarn1240 N43.68 E2.74 20.4.97 hare

21/98 Roc de Montalet Tarn1240 N43.68 E2.74 20.4.98 hare

22/98 Bois de Lause Tarn1200 N43.67 E2.8720.4.98 mouflon

23/98 Soumal,Douch Herault 950 N43.62 E2.9722.4.98 mouflon

24/98 Soumal,Douch Herault 950 N43.62 E2.9722.4.98 mouflon

25/98 Salis,Douch Herault750 N43.62 E2.95 22.4.98 hare

26/98 Douch Herault 920 N43.62 E2.9722.4.98 hare

50/98 Garabit Cantal850 N44.97 E3.18 8.6.98 rabbit

51/98 Montagne de Herault 500 N43.63 E3.37 11.6.98 hare Liausson

52/98 Monts de Herault 900 N43.67 E2.93 16.6.98 cattle l’Espinouse

53/98 Monts de Herault 900 N43.67 E2.95 16.6.98 hare l’Espinouse

54/98 Monts de Herault 900 N43.67 E2.93 16.6.98 deer l’Espinouse 160 M.J. Richardson

Table 1. Details of French dung samples and collection localities (continued)

Sample Elevation Latitude Longitude Locality Département Date Substrate no.* (m a.s.l.) (°) (°)

55/98 le Caylar Herault 800 N43.89 E3.30 18.6.98 sheep

56/98 le Caylar Herault 800 N43.89 E3.30 18.6.98 sheep

57/98 le Caylar Herault 800 N43.89 E3.30 18.6.98 rabbit

13/99 Gourdon Lot300 N44.68 E1.423.6.99 sheep

15/99 Carnac Morbihan15 N47.60 W3.0814.6.99 horse

23/99 le Fourquet Lot300 N44.73 E1.40 4.8.99 deer

3/01 Île Cimetiere Kerguelen 40 S49.49 E70.08 31.1.01 rabbit

4/01 Port-aux-Francais Kerguelen 100 S49.35 E70.22 13.2.01 rabbit

5/01 Col de SerraCorsica 360 N42.97 E9.37 9.4.01goat

6/01 Cascades des Corsica 1100 N42.12 E9.10 10.4.01 sheep Anglais

7/01 Acqua DoriaCorsica 250 N41.76 E8.72 12.4.01 sheep

8/01 S of Col de LavaCorsica 400 N42.23 E8.6413.4.01goat

9/01 OtaCorsica 350 N42.26 E8.76 13.4.01donkey

11/01 Île Cimetiere Kerguelen 40 S49.49 E70.084.1.01 rabbit

12/01 Île Cimetiere Kerguelen 40 S49.49 E70.084.1.01 rabbit

13/01 Île Cimetiere Kerguelen 40 S49.49 E70.084.1.01 rabbit

14/01 Île Cimetiere Kerguelen 40 S49.49 E70.084.1.01 rabbit

15/01 Île Haute Kerguelen 50 S49.39 E69.936.2.01 sheep

16/01 Île Haute Kerguelen 50 S49.39 E69.936.2.01 sheep

17/01 Île Haute Kerguelen 50 S49.39 E69.936.2.01 sheep

86/01 Lac de Montbel, Ariège 400 N42.96 E2.02 4.11.01horse Puivert

03/02 Espéraza cross, Aude 340 N42.94 E2.22 28.3.02 rabbit Aude

04/02 W of Espéraza Aude 350 N42.94 E2.22 28.3.02 sheep cross

05/02 W of Espéraza Aude 350 N42.94 E2.22 28.3.02 deer cross French coprophilous fungi 161

Table 1. Details of French dung samples and collection localities (continued)

Sample Elevation Latitude Longitude Locality Département Date Substrate no.* (m a.s.l.) (°) (°)

06/02 Route D54,Arques Aude 460 N42.96 E2.381.4.02 hare

07/02 les Plas,Antugnac Aude 370 N42.94 E2.23 5.4.02 hare

08/02 Camps de la Borde, Aude 400 N42.85 E2.22 6.4.02 deer Cavirac

09/02 Bitrague,Quillan Aude 300 N42.88 E2.19 6.4.02 rabbit

17/02 Pic de Brau, Aude 600 N43.01 E2.23 1.7.02 sheep Roquetaillade

18/02 Pic de Brau, Aude 620 N43.01 E2.23 1.7.02 sheep Roquetaillade

19/02 Pic de Brau, Aude 620 N43.01 E2.23 1.7.02 hare Roquetaillade

08/03 Soumal,Douch Herault 940 N43.61 E2.96 18.3.03 mouflon

09/03 Soumal,Douch Herault 940 N43.61 E2.96 18.3.03 hare

10/03 Soumal,Douch Herault 960 N43.61 E2.97 18.3.03 rabbit

11/03 path to Pic de Aude 800 N42.87 E2.38 21.3.03 cattle Bugarach

12/03 la Frau Haute Aude 600 N42.97 E2.36 23.3.03 horse

56/03 Quéribus, Aude 640 N42.84 E2.62 15.11.03 goat Cucugnan

57/03 road to le Poux, Aude 300 N42.94 E2.2117.11.03 rabbit Espéraza

18/04 Col de Jau Aude 1510 N42.69 E2.25 21.4.04hare

19/04 Pic Dourmidou Aude/ 1840 N42.71 E2.26 21.4.04hare Pyrénées- Orientales

20/04 Pic Dourmidou Aude/ 1840 N42.71 E2.26 21.4.04 cattle Pyrénées- Orientales

21/04 Col de Jau Aude 1510 N42.69 E2.25 21.4.04 vole?

22/04 -le-Château Aude 400 N42.92 E2.26 24.4.04hare

88/04 Granès,Quillan Aude 300 N42.89 E2.2111.9.04hare

89/04 Antugnac, CouizaAude 470 N42.95 E2.2112.9.04hare 162 M.J. Richardson

Table 1. Details of French dung samples and collection localities (continued)

Sample Elevation Latitude Longitude Locality Département Date Substrate no.* (m a.s.l.) (°) (°)

1/05 Domaine de Aude 250 N42.95 E2.23 14.3.05hare Garenaud, Montazels

28/05near Fa Aude 400 N42.93 E2.20 21.6.05hare

29/05 GR36,Col de Aude 575 N42.87 E2.54 22.6.05 cattle Gardiole, nr Peyrepertuse

30/05 GR36,Col de Aude 565 N42.87 E2.55 22.6.05 sheep Gardiole, nr Peyrepertuse

31/05 La Girude, Aude 460 N42.93 E2.20 22.6.05 sheep/goat Soulatgé

6/06 nr la Toupine, Aude 420 N42.91 E2.22 28.4.06 hare St-Ferriol

1/07 Fond Heliot, Guadeloupe 20 N16.28 W61.80 8.2.07 goat Deshaies, Basse- Terre

2/07 Fond Heliot, Guadeloupe 20 N16.28 W61.80 8.2.07 donkey Deshaies, Basse- Terre

3/07 Roches Caraibes, Guadeloupe 200 N16.03 W61.748.2.07 cattle? Bailiff, Basse-Terre

4/07 nr Plage Bois Jolan, Guadeloupe 5 N16.23 W61.3511.2.07 goat St-Anne,Grande- Terre

5/07 nr Plage Bois Jolan, Guadeloupe 5 N16.23 W61.3511.2.07 cattle St-Anne,Grande- Terre

44/07 Tourtrès -et- 175 N44.51 E0.43 11.10.07 rabbit Garonne

45/07 D619, road junction Pyrénées- 880 N42.67 E2.43 15.10.07 hare to Comes,Eus Orientales

46/07 Camp Gast,Puivert Aude 610 N42.90 E2.07 21.10.07 cattle

50/07 la Coume,EspérazaAude 450 N42.93 E2.20 27.12.07 hare

* = MJR sample no. and year identifier French coprophilous fungi 163 curve was plotted for the samples from mainland France and Corsica. The equation for that curve was calculated (y = axb, where y = cumulative no. of species observed in x samples) and solved for x = 50 samples. That value was compared with those obtained from the same assemblage of species recorded in a worldwide study of a similar range of substrates (Richardson 2001a).

RESULTS

The records are listed by ‘département’ in alphabetical order, and within each département in order of collection. They are treated in two ways; common and ubiquitous species are simply listed, with the sample number/year and département from which they were recorded. More interuestnusuinga olr records are treated in more detail, with descriptions and discussion in the context of information from observations from over 1000 other samples collected worldwide, yielding over 10000 records of coprophilous fungi. Details of incompletely identified taxa are provided for future reference. Dried material and/or slides for collections marked E have been placed in the Herbarium of the Royal Botanic Garden,Edinburgh. ZYGOMYCOTINA Cunninghamella echinulata (Thaxt.) Thaxt. ex Blakeslee Guadeloupe: 5/07 Mortierella bainieri Costantin Kerguelen: 14/01 Phycomyces nitens Kunze Herault: 117/97 Pilaira anomala (Ces.) J. Schröt. Aude: 3/02, 8/02, 9/02, 1/05; Herault: 117/97, 26/98 Pilaira moreaui Y. Ling Herault: 23/98, 10/03; Lot-et-Garonne: 44/07 Pilobolus crystallinus . crystallinus (F. H. Wigg.) Tode Aude: 5/02, 18/02, 56/03, 1/05, 30/05, 31/05; Corsica: 5/01; Guadeloupe: 1/07; Lot: 13/99, 23/99 Pilobolus crystallinus var. kleinii (Tiegh.) R.Y. Zheng &G.Q. Cheng Aude: 30/05, 46/07; Corsica: 7/01; Guadeloupe: 2/07, 4/07; Kerguelen: 16/01, 17/01 Pilobolus roridus var. umbonatus (Buller) F.M. Hu &R.Y. Zheng Aude: 18/02; Herault: 106/97, 109/97, 115/97, 23/98, 24/98, 8/03, 10/03; : 22/98 Piptocephalis lepidula (Marchal) R. K. Benj. Herault: 25/98 Members of the genus are of interest because of their obligate parasitism on other Mucorales, and they are quite frequently recorded in the early stages of incubation of dung samples, when host species are most frequently seen. P. lepidula is recognised by its dry heads with two-celled merosporangia, and fusoid spores 4-5 × 2-3 µm. ASCOMYCOTINA Ascobolus albidus Crouan Aude: 12/03; Herault: 115/97, 24/98, 26/98, 54/98; Tarn: 22/98 164 M.J. Richardson

Ascobolus cervinus Berk. &Broome Herault: 106/07, 115/97 Ascobolus hawaiiensis Brumm. Aude: 4/02, 5/02; Herault: 110/97, 56/98; Pyrénées-Orientales: 45/07 A coprophil with a worldwide distribution, in spite of its specific name. It has now been recorded from many countries, from Iceland, UK and Greece in the northern hemisphere through to Chile,Australia, New Zealand and the Falkland Islands in the southern hemisphere (Richardson 2004). Ascobolus immersus Pers. Aude: 5/02, 18E/02, 28/05, 29/05, 30/05, 46/07, 50/07; Aveyron: 112/97; Corsica: 9/ 01; Herault: 110/97, 114/97, 115/97, 23/98, 24/98, 52/98, 55/98; Guadeloupe: 2/07, 3/ 07, 4/07, 5/07; Pyrénées-Orientales: 45/07 Ascobolus sacchariferus Brumm. Aude: 8/02; Corsica: 5E/01; Herault: 24/98; Tarn: 18E/98, 22E/98 Described from the Netherlands by van Brummelen (1967) from two samples of deer dung, the substrate range of this species is increased by its occurrence also on sheep, hare and goat dung among the French samples, and I also have a record on rabbit from Scotland. Apart from the substrate differences, the collections agree well with van Brummelen’s original description, with characteristically white, superficial apothecia with a granular excipular surface. fimicola (de Not. & Bagl.) Dennis Aude: 11/03 Coprotus cf. disculus Kimbr.,Luck-Allen & Cain Aude: 4/02, 8/02 Coprotus granuliformis (P. Crouan &H. Crouan) Kimbr. Aude: 11/03, 30/05 Coprotus sexdecimsporus (P. Crouan &H. Crouan) Kimbr. Aude: 5/02, 8/02, 22E/04, 46/07, 50/07; Aude/Pyrénées-Orientales: 20/04; Aveyron: 112/97; Herault: 110/97, 114E/97, 55/98; Kerguelen: 13/99, 15/99; Pyrénées- Orientales: 45/07 Coprotus spp. Aude: 17/02, 1E/05, 29/05; Aude/Pyrénées-Orientales: 20/04; Corsica: 8/01, 9/01 Without a modern monograph of the genus, the identification of small white Coprotus species with 8-spored asci presents difficulties. The morphological features of these 6 collections, particularly ascus size and shape and spore size and arrangement, are sufficiently distinct to allow the six to be considered different from each other, but do not agree well with the description of existing species. Indeed one (9/01) produced apothecia with dimorphic asci and spores – asci 95-100 × 26-29 µm and 50 × 8-10 µm, and spores 16 × 9-10 µm and 6 × 3 µm. – perhaps a ‘chimera’ composed of two species forming a single apothecium. Iodophanus carneus (Pers.) Korf Aude: 4/02, 17E/02, 18/02, 19/02, 56E/03, 88/04, 1/05, 30/05, 46E/07; Aveyron: 113/97; Corsica: 7E/01, 9/01; Guadeloupe: 1/07, 4/07; Herault: 114/97, 55/98, 56/98; Kerguelen: 4/01,11/01, 14/01; Lot: 23/99 Lasiobolus ciliatus (J. C. Schmidt:Fr.) Boud. Corsica: 6/01; Herault: 51/98; Lot: 13/99 Lasiobolus cuniculi Velen. Ariège: 86/01; Aude: 4/02, 6/02, 29/05, 46/07; Corsica: 5/01, 8/01, 9/01; Herault: 107/97, 108/97, 115/97, 24/98, 52/98, 55/98 Lasiobolus diversisporus (Fuckel) Sacc. Aude: 11/03, 12/03; Pyrénées-Orientales: 45/07 French coprophilous fungi 165

Lasiobolus ruber (Quél.) Sacc. Aude: 12/03 Peziza bovina Phill. Corsica: 9/01 Peziza fimeti (Fuckel) Seaver Aude: 29E/05 Peziza vesiculosa Bull. Herault: 55E/98 Saccobolus cf. beckii Heimerl Aude: 12/03 Distinguished from S. versicolor by the very thick and warted nature of the epispore. I observe a range of spore ornamentation, from smooth, through lightly cracked or flaky to quite rough, often from the same apothecium, in collections that I am content to identify as S. versicolor. The spores of this collection were on occasion notably very rough, although not consistently so, and are with hesitation identified as near to S. beckii. Saccobolus citrinus Boud. &Torrend Aude: 46/07; Corsica: 7/01, 9/01; Guadeloupe: 2/07, 3/07, 4/07, 5/07; Lot: 13/99, 23/99 Saccobolus depauperatus (Berk. &Broome) E.C. Hansen Aude: 5/02; Corsica: 7/01, 8/01; Herault: 51/98; Lot: 13/99, 23/99; : 15E/99 Saccobolus truncatus Vel. Guadeloupe: 2/07, 3/07, 4/07, 5E/07 Van Brummelen (1967) observes that this is a widely distributed species, but in my experience it tends to occur at lower latitudes, with all but one of twelve other records from samples collected between 31°N and 20°S, so its occurrence on French samples only in the Caribbean region is perhaps not surprising. Saccobolus verrucisporus Brumm. Guadeloupe: 4/07 Saccobolus versicolor (P. Karst.) P. Karst. Aude: 5/02, 88/04, 1/05, 30E/05; Aude/Pyrénées-Orientales: 20/04; Aveyron: 113/ 97; : 50/98; Herault: 114/97, 117/97, 52/98, 57/98; Pyrénées-Orientales: 45/07 Thecotheus holmskjoldii (E. C. Hansen) Chenant. Aude: 4E/02, 8/02, 17/02, 18E/02; Corsica: 9/01 Thecotheus species are widespread, but not very common. I have 23 records of the genus from a total of over 10000 coprophil records, twelve of which are of T. holmskjoldii, on which basis it is one of the commonest species of the genus. Aas (1992) cites numerous collections worldwide, and examined material collected by Lundqvist from the Alpes de Haute Provence (Gorge du Verdon), and two collections from Corsica. Thecotheus lundqvistii Aas Aude/Pyrénées-Orientales: 20E/04 Thecotheus apothecia often take some time to appear in moist chamber culture, but after just 8 d’incubation a pure white, concave apothecium, 2.5 mm diam. was observed on a sample of cattle dung. The spores of the French collection were obliquely uniseriate, ellipsoid, symmetrical, 22.5-24 (-25.5) × 12.5- 13 µm, smooth, with a hemispherical papilla at each pole, and a distinct gel. It is interesting that the sample was collected from the summit of a high pasture hill (elevation 1840 m) in the foothills of the Pyrénées, still with large snow patches and no cattle on the hill, so it is assumed that the dung was deposited the previous autumn and would have passed much of the winter under snow cover. It is apparently rare, with records only from Sweden and Spain (Aas, 1992),Italy (Doveri, 2004), and Greece (Richardson, 2008). 166 M.J. Richardson

Trichobolus sphaerosporus Kimbr. Corsica: 5/01; Herault: 54/98, and Trichobolus zukalii (Heimerl) Kimbr. Aude: 8/02, 56/03, 31/05; Corsica: 8/01 Trichobolus species occur early in the incubation period. They are infrequently recorded, although Doveri (2004) has described T. zukalii from 29 Italian collections, and discussed the differences between T. zukalii and T. sphaerosporus. Doveri distinguishes the two species particularly on the basis of spore morphology, considering collections with globose to subglobose spores with a Q (l/w) value of 1-1.16 (mean = 1.07), and with smaller spores (8.9-9.9 × 8.4- 9.4 µm) to be T. sphaerosporus, and those with subglobose to predominantly ellipsoid spores (Q = 1-1.26, mean 1.19), and larger spores (10.5-12.6 × 9.4-10.5 µm) to be T. zukalii. Dissing (in Hansen &Knudsen 2000) distinguishes the two species on the basis of presence (T. zukalii) or absence (T. sphaerosporus) of de Bary bubbles in the spores, and gives very similar sizes for the spores of both species (T. zukalii: 9-11 × 7-8.5 µm, T. sphaerosporus: 9-10.5 × 8.5-9 µm), describing them both as subglobose. Doveri (2004), conscious of the readiness with which artefacts can be formed, prefers not to place reliance on the presence or absence of de Bary bubbles as a diagnostic feature, and I rarely observe them. Given the variation in interpretation of these two species, and the variation observed in material from individual collections,I treat the six French collections together. I have examined the spore sizes from 25 collections, including the 6 French ones, with the results as illustrated in Fig. 2, in which the minimum and maximum spore lengths and widths are plotted. There is considerable overlap between those identified as T. sphaerosporus (mean 10.4 × 9.6 µm,Q = 1.0-1.13 (mean = 1.08) and T. zukalii (mean 9.7 × 8.3 µm,Q = 1.14-1.21 (mean = 1.17), and it is still open to debate as to whether or not these differences are sufficient to separate two species, or represent the opposite ends of the range of a single species, one end with a tendency to have larger more rounded spores, the other with smaller more ellipsoid spores. These values agree very well with those given by Doveri (2004) for the material he has studied. In the French material apothecia were up to 350 µm diam., immersed, setose, each with a single polysporus ascus. In 54/98 setae were up to 420 µm long, hyaline, thick-walled, with septa 20-30 µm apart; ascospores hyaline, globose to broad ellipsoid, 9.5-10 × 9-9.5 µm. In 5/01 setae were shorter, up to 225 µm long and spores were a little larger, but still almost globose to broad ellipsoid, 9-12 × 8–11 µm. These two collections were determined as T. sphaerosporus. The other four collections, with sub-globose to ellipsoid spores 8-11 × 8-9.5 µm, were determined as T. zukalii.

Thelebolales Ascozonus woolhopensis (Renny) Boud. Aude: 3E/02, 6/02, 9/02; Tarn: 18E/98 Thelebolus microsporus (Berk. &Broome) Kimbr. Herault: 24/98 Thelebolus stercoreus Tode Aude: 3E/02, 6/02, 7/02, 1/05; Herault: 106/97, 107/97, 109/97, 24/98, 25/98, 26/98, 51/98, 53/98, 9/03, 10/03; Tarn: 18E/98, 20/98, 21/98, 22E/98 De Hoog et al. (2005), on the basis of molecular studies, accept only four species of Thelebolus: T. stercoreus, T. microsporus, and two new species described from biomats in Antarctica. They found that many cultures from phenotypically very different teleomorphs, including cultures from uniascal and polyascal types, with small to large asci, and few to very many-spored types, and French coprophilous fungi 167

13

12

11 m) µ

10

9

8 Smaller spore measurement (

7

6 7 8 9 10 11 12 13 Larger spore measurement (µm) Fig. 2. Plots to show range of spore size recorded for 11 collections of Trichobolus sphaerosporus (unbroken lines, mean = ●) and 14 collections of T. zukalii (broken lines, mean = ▲). The straight line represents ‘length = width’, i.e. spores spherical; points below that line indicate a tendency towards spores being ellipsoid.

with many names, are molecularly indistinguishable from each other and from cultures of T. stercoreus from material with the classical morphological criteria of a single large ascus with 2000+ spores, and ecological preference for lagomorph dung. The records associated with the T. stercoreus entry above refer to that morphological understanding of T. stercoreus, and it was present on 13 of the 24 French samples of hare dung. The following, according to the above, should be considered as forms of T. stercoreus, but were originally recorded as: Thelebolus caninus (Auersw.) Jeng &J.C. Krug Aude: 1/05 Thelebolus crustaceus (Fuckel) Kimbr. Herault: 21/98, 53/98 Thelebolus nanus Heimerl Aude: 5/02; Herault: 109/97, 25/98; Lot-et-Garonne: 44/07; Pyrénées-Orientales: 45/07 168 M.J. Richardson

Thelebolus polysporus (P. Karst.) Otani & Kanzawa Aude: 3/02, 4/02, 6/02, 7/02, 1/05, 50/07; Cantal: 50/98; Herault: 107/97, 26/98, 52/98, 57/98, 9/03; Kerguelen: 3/01, 4/01, 11/01, 12/01, 14/01; Lot-et-Garonne: 44/07 Thelebolus sp. Aude: 56/03; Aude/Pyrénées-Orientales: 19/04; Herault: 25/98

Helotiales Orbilia leporina Velen. Aude: 11/03, 12/03; Herault: 116/97; Pyrénées-Orientales: 45/07

Sordariales Anopodium ampullaceum N. Lundq. Herault: 109E/97; Tarn: 20E/98 Details of one of the French collections (109/97) were included in a discussion by Richardson (1998) of the various names given to the taxon, which has pedicellate spores like Podospora and Schizothecium but is unusual in having the pedicel directed towards the apex of the ascus. Since then another French collection has been found (20/98). This would seem to be a temperate species, recorded also from the UK and Sweden (unpublished), and from Sweden,Norway and Belgium by Lundqvist (1972), and the two French records would seem to be the most southerly so far. It occurs mainly on lagomorph dung (34 records), with occasional occurrences on other substrates (2 vole, 1 deer and 1 goose). Apodospora gotlandica N. Lundq. Aude: 29E/05 There appear to be no reports of this since its description by Lundqvist (1972) from Swedish horse dung collected in 1959, so the French collection is described. Perithecia large, up to 750 µm diam, reddish brown below and opaque, so difficult to make out any detailed cellular structure, with a very dark, short (110 µm), opaque neck. Asci 8-spored, 320-385 × 23-27 µm, cylindrical above, tapering below to 100 µm long stipe, and with a marked KI-ve apical pore. Spores uniseriate, smooth, blackish brown, ellipsoid, (32-) 38-42 × (16-) 17.5- 21 µm, with a faint germ pore at the distal end. Gel broad and uniform, expanding to 10 µm in water, and with an apical invagination. Arnium arizonense (Griffiths) N. Lundq. &J.C. Krug Aude: 4E/02, 5E/02, 18E/02, 30E/05, 31/05, 46/07; Corsica: 9/01; Lot: 23/99 Arnium is distinguished from related genera by the possession of gelatinous appendages on the ascospores, but no pedicel or primary appendage. A. arizonense is so far unique in having 4-spored asci. French collections had clavate asci 290-340 × 32-38 µm, and spores 41.5-55 × 19-26 µm, with tapering gelatinous appendages up to 200 µm long × 8-13 µm wide at their base, inserted asymmetrically at each end of the spore. Arnium caballinum N. Lundq. Aude: 12/03, 29E/05; Corsica: 6/01 Arnium hirtum (E.C. Hansen) N. Lundq. &J. C. Krug Aude: 30E/05; Aveyron: 113E/97; Herault: 108E/97 Arnium imitans N. Lundq. Aude: 29E/05 Arnium leporinum (Cain) N. Lundq. &J. C. Krug Herault: 108E/97 Arnium mendax N. Lundq. Aude: 11E/03; Corsica: 8E/01; Herault: 115E /97 French coprophilous fungi 169

Arnium sp. Aude/Pyrénées-Orientales: 20/04 Positive identification was not possible, with only limited material seen. The perithecium was schizothecioid in structure, the ascus form typical of an Arnium sp., and with no apical structure, and spores very small for 8-spored Arnium species, 19-19.5 × 12 µm. Bombardioidea stercoris (DC.) N. Lundq. Herault: 51E/98 Cercophora cf. anisura N. Lundq. Aude: 29E/05 A tentative identification of limited and immature material, based on the schizothecioid structure of the perithecium, vermifom spores 42-52 × 4.5 µm, with appendages of unequal length. Cercophora sordarioides (Speg.) N. Lundq. Aude: 12E/03 Chaetomium bostrychodes Zopf Aude: 9/02; Cantal: 50/98; Herault: 109/97, 51/98; Kerguelen: 13/01, 14/01; Lot-et- Garonne: 44/07 Chaetomium cf. cuniculorum Fuckel Aude: 1/05 Coniochaeta hansenii (Oudem.) Cain Aude: 6E/02; Aude/Pyrénées-Orientales: 19/04; Herault: 117E/97, 53/98 Coniochaeta leucoplaca (Berk. & Ravenel) Cain Aude: 11/03, 22/04; Aude/Pyrénées-Orientales: 20E/04; Cantal: 50E/98 Coniochaeta ligniaria (Grev.) Massee Aude: 17/02, 18E/04, 21E/04; Aude/Pyrénées-Orientales: 19E/04; Aveyron: 113/97; Corsica: 8/01; Herault: 109/97, 26/98, 52/98, 56/98, 9E/03; Pyrénées-Orientales: 45/07; Tarn: 19/98, 20/98, 21/98 Coniochaeta scatigena (Berk. &Broome) Cain Aude: 18E/04, 29/05; Corsica: 6/01; Herault: 116/97,117/97, 52/98, 53/98, 54/98, 9E/03 Fimetariella microsperma J.C. Krug &J.H. Mirza Herault: 114/97 Podospora australis (Speg.) Niessl Aude: 4E/02, 5E/02, 18/02 Podospora bifida N. Lundq. Herault: 115/97 Podospora communis (Speg.) Niessl Guadeloupe: 1E/07, 2/07, 5/07 This is one of three species that appears to be characteristic of the per- ithecial mycobiota of dung from lower latitudes, with P. immersa and P. longi- caudata (q.v.)(Richardson, in press). P. communis is the most frequent of the three and, although Lundqvist (1972) noted that it ‘seems to be worldwide, … records from the tropics are still very few’, and he refers to unverified records from France, my experience is that it is more frequent in samples from lower lat- itudes, with 24 of 30 of my collections from within the tropics. Podospora curvicolla (G. Winter) Niessl Ariège: 86/01 Podospora decipiens (G. Winter ex Fuckel) Niessl Ariège: 86E/01; Aude: 4/02, 6/02, 57/03, 22E/04, 29/05, 30E/05, 31/05, 46/07; Aude/ Pyrénées-Orientales: 20/04; Aveyron: 112/97, 113/97; Corsica: 6/01, 9/01; Herault: 170 M.J. Richardson

110/97, 114/97, 115/97, 116/97, 24E/98, 52/98, 55/98; Lot 13/99, 23/99; Pyrénées- Orientales: 45/07; Tarn: 19/98, 20/98, Podospora gigantea Mirza & Cain Corsica: 9E/01 Podospora gwynne-vaughaniae (Page) Cain Herault: 115/97 Podospora immersa (R. Stratton) Cain Guadeloupe: 1E/07, 5/07 A lower latitude species with, in addition to these two records from Guadeloupe, six other occurrences from the Caribbean region (Richardson, in press), none from temperate regions in my collections or studied by Lundqvist (1972), and the type and other collections from the USA and Mexico all from south of 40°N (Mirza & Cain, 1969). Podospora intestinacea N. Lundq. Aude: 11E/03, 12E/03, 29E/05; Corsica: 6E/01, 9/01 Podospora longicaudata (Griffiths) Cain Guadeloupe: 2/07, 3E/07, 4/07 A lower latitude species, with three records from Guadeloupe, three from St Lucia (Richardson, in press), one from Brazil (Richardson 2001b), none from temperate regions in my collections or studied by Lundqvist (1972), and the type and other collections from the USA, Puerto Rico,Mexico and Pakistan all from south of 40°N (Mirza & Cain, 1969). Podospora myriaspora (P. Crouan &H. Crouan) Niessl Aude: 28/05; Herault: 52/98 Podospora pauciseta (Ces.) Traverso Aude: 31/05; Corsica: 9/01; Guadeloupe: 2/07, 4/07, 5/07; Lot: 13/99, 23/99; Morbihan: 15/99 Podospora pleiospora (G. Winter) Niessl Aude: 8/02, 18/02, 22/04, 89/04, 1/05; Aveyron: 113/97; Cantal: 50/98; Herault: 110/ 97, 115/97, 24E/98, 25/98, 26/98, 51E/98, 55/98, 57/98; Lot 13/99, 23/99; Lot-et- Garonne: 44/07; Pyrénées-Orientales: 45/07 Podospora pyriformis (Bayer) Cain Corsica: 8E/01 Podospora setosa (G. Winter) Niessl Aude: 8E/02, 31/05; Corsica: 7/01; Lot: 13/99, 23/99; Lot-et-Garonne: 44/07 Podospora cf. similis (E.C.Hansen) Niessl Aude/Pyrénées-Orientales: 20/04; Corsica: 6E/01 There are relatively few species of Podospora/Schizothecium with 16-spored asci, and details of the collections are given as they differ from those described. Perithecia dark, opaque, pyriform, 525-900 µm high × 200-410 µm diam, flexuose hairy below and distinctly setose at the neck, with dark septate hyphae, paler towards the tip, <200 µm long, some aggregated into asymmetrically arranged fascicles around the neck. Asci 16-spored, 225-340 × 35-70 µm, fusoid, tapering below, with no apical apparatus. Spores ellipsoid, the dark cell 32-35 × 16-19 µm, with apical germ pore. Pedicel small, 6-10 × 2-3 µm. Secondary appendages not obvious, but the whole spore is surrounded by a faint halo when mounted in Indian ink. Schizothecium dubium has spores of similar size, but is distinctly schizothecioid in perithecial structure, without rigid hairs, and has persistent appendages; P. pleiospora has characteristic sect. Rhypophila spores and tubercules at the perithecial neck, which were absent from this material. They are nearest to P. similis, but that is described without rigid setae, and with spore appendages (Mirza & Cain 1969), so this may be an undescribed species. French coprophilous fungi 171

Schizothecium conicum (Fuckel) N. Lundq. Aude: 8/02, 11/03, 12/03, 21E/04, 28/05, 29/05, 30/05, 31/05; Aude/Pyrénées- Orientales: 19/04, 20/04; Herault: 114/97, 52/98; Kerguelen: 12/01; Lot: 23/99 Schizothecium dakotense (Griffiths) N. Lundq. Aude: 57E/03; Cantal: 50E/98 Schizothecium miniglutinans (Mirza & Cain) N. Lundq. Herault: 9/03, 10E/03; Pyrénées-Orientales: 45/07 Schizothecium tetrasporum (G. Winter) N. Lundq. Aude: 3/02, 4/02, 6/02, 7E/02, 8/02, 9/02, 19/02, 57/03, 21E/04, 88E/04, 1/05; Aveyron: 113/97; Cantal: 50/98; Herault: 108/97, 109/97, 117/97, 24/98, 26/98, 54/98, 57/98, 8/03, 9/03, 10E/03; Kerguelen: 4/01; Lot-et-Garonne: 44/07; Tarn: 18/98 Schizothecium vesticola (Berk. &Broome) N. Lundq. Ariège: 86/01; Aude: 4/02, 5/02, 8/02, 18/02, 28/05, 29/05, 31/05, 46/07, 50/07; Aveyron: 113/97; Cantal: 50/98; Corsica: 5/01, 6/01, 7/01, 8/01, 9/01; Herault: 108/97, 109/97, 110/97, 115/97, 116E/97, 117/97, 24/98, 25/98, 26/98, 52/98, 53/98, 54/98, 55/ 98, 8/03; Lot: 13/99, 23/99; Pyrénées-Orientales: 45/07 Sordaria alcina N. Lundq. Herault: 23E/98, 25/98, 8/03; Tarn: 18/98, 22E/98 Sordaria fimicola (Roberge ex. Desm.) Ces. & de Not. Ariège: 86E/01; Aude: 18/02, 19/02, 56/03, 88/04, 89/04, 1/05, 28/05, 6E/06; Aveyron: 111/97; Cantal: 50/98; Corsica: 5/01; Herault: 106/97, 107/97, 108/97, 109/97, 115/97, 117/97, 51/98, 52/98, 53/98, 57/98, 8/03, 9/03; Lot: 13/99; Morbihan: 15/99; Tarn: 18/98 Sordaria humana (Fuckel) G. Winter Aude: 18/04, 1/05; Herault: 106/97,107/97, 23/98, 26/98, 52/98, 53/98, 9/03; Lot: 13/ 99; Morbihan: 15/99 Sordaria macrospora Auersw. Aude: 89E/04 Sordaria superba de Not. Cantal: 50E/98; Tarn: 18/98 Zygopleurage zygospora (Speg.) Boedijn Guadeloupe: 5/07 Zygospermella insignis (Mouton) Cain Aude: 11E/03, 29E/05; Aude/Pyrénées-Orientales: 20E/04 Hypocreales Melanospora fusispora (Petch) Doguet Herault: 24/98, 25E/98 Selinia pulchra (G. Winter) Sacc. Aude: 5E/02; Herault: 114E/97 Microascales Cephalotrichum stemonitis (Pers.) Nees Kerguelen: 3/01, 4/01, 11/01, 13/01, 14/01, 15/01 Occurring as the anamorphic form, Doratomyces stemonitis (Pers.) F.J. Morton &G. Sm. Noticeably prevalent on the samples from the Kerguelen Islands, perhaps favoured by the conditions in which they had been kept before or during transmission, since it usually only develops occasionally in my cultures. Viennotidia fimicola (Marchal) P. F. Cannon &D. Hawksw. Herault: 115/97 172 M.J. Richardson

Onygenales Arachniotus ruber (Tiegh.) J. Schröt. Kerguelen: 16E/01 Gymnoascus reessii Baran. Aude: 9E/02; Herault: 25E/98

Xylariales Hypocopra brefeldii Zopf Aude: 22E/04, 89/04, 1/05, 28/05, 6E/06, 50/07; Aveyron: 113/97; Herault: 26/98, 51E/98; Pyrénées-Orientales: 45/07 Hypocopra equorum (Fuckel) G. Winter Aveyron: 111/97 Hypocopra festucacea J. C. Krug & Cain Herault: 56E/98 Hypocopra merdaria (Fr.) Kickx f. Aude: 17/02 Hypocopra ornithophila Speg. Aude/Pyrénées-Orientales: 19/04 Hypocopra parvula Griffiths Aude: 12E/03 For a discussion of the identity of this material see Richardson (2004). Krug (in litt.) has suggested that the European material determined as H. parvula is a different species still to be described. Hypocopra stephanophora J. C. Krug & Cain Herault: 107E/97, 117E/97 Hypocopra stercoraria (Sowerby) Sacc. Herault: 56E/98 Phomatospora minutissima (P. Crouan &H. Crouan) N. Lundq. Guadeloupe: 4/07, 5E/07; Herault: 114E/97; Lot: 23/99 This fungus is probably commoner than records indicate. It is usually seen only after a long period of incubation (at least 1 month, and often much longer, Richardson, 2002). The perithecia are very small, <150 µm diam., and immersed in the substrate, with only the erumpent neck and ostiole visible. P. minutissima has obliquely uniseriate, ellipsoid hyaline spores, 5-6 × 3 µm, while the very similar P. coprophila M. J. Richardson has spores that are short-rod shaped aligned linearly in the ascus. P. coprophila is frequent in the UK, and I have a record from the Falkland Islands; P. minutissima is widespread, and in addition to the French collections I have records from Brazil,Australia, St Helena, St Lucia and Dominica. Both species seem to occur most frequently on the dung of ruminants. Podosordaria tulasnei (Nitschke) Dennis Aude: 88/04, 30/05, 6/06; Aveyron: 111/97, 113/97; Herault: 106/97, 107/97, 108/97, 110/97, 24/98, 25/98, 26/98, 52/98, 53/98, 55/98, 57/98; Kerguelen: 4/01; Tarn: 18/98, 22/98 Identification of this species is based on the development of the characteristic long, branching stromata that grow from the dung. They have never produced perithecia in my cultures.

Dothideales Delitschia excentrica Griffiths Aude: 17E/02 Delitschia furfuracea Niessl Aude: 57E/03; Herault: 107/97, 109E/97, 24/98, 25E/98 French coprophilous fungi 173

Delitschia marchalii Berl. &Voglino Aude/Pyrénées-Orientales: 19/04 Delitschia niesslii Oudem. Aude: 22E/04 Delitschia patagonica Speg. Aude: 18/04, 50/07; Herault: 9E/03 Delitschia perpusilla Speg. Aude: 22/04; Herault: 56/98 Delitschia tomentosa Luck-Allen & Cain Aude/Pyrénées-Orientales: 19/04 Delitschia winteri Phill. &Plowright Ariège: 86/01; Aude: 4/02, 46/07; Aude/Pyrénées-Orientales: 20/04; Aveyron: 112/97; Corsica: 9/01; Herault: 114/97, 117E/97 Preussia funiculata (Preuss) Fuckel Pyrénées-Orientales: 45/07 Sporormia fimetaria (De Not.) De Not. Herault: 108E/97, 115/97, 26E/98, 8/03 A widespread but infrequently recorded fungus, characterised by very small pseudothecia, ca 100 µm diam, and the eight 16-celled spores amalgamated into a bundle, resembling a maize cob, with terminal gelatinous appendages. Recorded from the UK, France,Morocco,Australia, USA and New Zealand. Sporormiella australis (Speg.) S. I. Ahmed & Cain Ariège: 86/01; Aude: 4/02, 5/02, 6/02, 7/02, 8/02, 9/02, 57/03; Aude/Pyrénées- Orientales: 20E/04; Aveyron: 111/97, 112/97; Cantal: 50/98; Corsica: 5/01, 6/01, 7/01, 8/01, 9/01; Herault: 106/97, 107/97, 108/97, 109/97, 114/97, 115/97, 117/97, 24/98, 25/98, 26/98, 52/98, 53/98, 54/98, 55/98, 57/98, 8/03, 9/03, 10E/03; Kerguelen: 4/01; Lot-et-Garonne: 44/07; Tarn: 18/98, 20/98, 21/98, 22/98 One of the commonest and most widespread coprophilous fungi, along with S. intermedia. At the upper end of its spore range there is often overlap with the lower end of that of S. intermedia, but on most occasions it is possible to distinguish them with confidence. Sporormiella bipartis (Cain) S.I. Ahmed & Cain Aude: 1/05; Tarn: 18/98, 20/98 Collection 1/05 differed from typical S. bipartis in having spores broader than normal, 9-9.5 µm cf. the more typical 6-7 µm, and a less obvious tendency for spores to break into 4-celled halves. Sporormiella borealis (I. Egeland) J.C. Krug Aude: 29/05 Sporormiella dubia S.I. Ahmed & Cain Aude: 22/04 Sporormiella grandispora S.I. Ahmed & Cain Aude: 11/03, 29/05, 46/07 Sporormiella heptamera (Auersw.) S.I. Ahmed & Cain Cantal: 50/98 Sporormiella intermedia (Auersw.) S.I. Ahmed & Cain Ariège: 86/01; Aude: 3/02, 6/02, 7E/02, 8/02, 9/02, 19/02, 11/03, 12/03, 56/03, 57/03, 18E/04, 22/04, 88/04, 89/04, 1/05, 28/05, 29/05, 6E/06, 46/07, 50/07; Aude/Pyrénées- Orientales: 19E/04, 20E/04; Aveyron: 111/97, 112/97, 113/97; Cantal: 50/98; Corsica: 5E/01, 6/01, 8/01; Herault: 107/97, 109/97, 110/97, 115/97, 117/97, 24/98, 25/98, 26/98, 51/98, 52/98, 53/98, 54/98, 55/98, 56/98, 57/98, 9/03, 10/03; Kerguelen: 4/01; Pyrénées-Orientales: 45/07; Tarn: 18/98, 19/98, 20/98, 21/98 174 M.J. Richardson

Sporormiella lageniformis (Fuckel) S.I. Ahmed & Cain Ariège: 86/01; Aude: 6/02, 12/03, 46/07; Aveyron: 113/97; Corsica: 8/01; Herault: 115/97, 116/97 Sporormiella leporina (Niessl) S.I. Ahmed & Cain Aude: 8E/02 Sporormiella longispora (Cain) S.I. Ahmed & Cain Aude: 1E/05 Sporormiella megalospora (Auersw.) S.I. Ahmed & Cain Aude: 4/02, 8/02,17/02; Aveyron: 112/97; Herault: 56/98 Sporormiella minima (Auersw.) S.I. Ahmed & Cain Ariège: 86/01; Aude: 4/02, 18/02, 19/02, 28/05, 31/05, 46/07; Aveyron: 111/97, 112/ 97; Guadeloupe: 3/07, 4/07; Herault: 109/97, 110/97, 114/97, 115/97, 117/97; Lot: 13/99 Sporormiella octonalis S.I. Ahmed & Cain Corsica: 6/01 Sporormiella ontariensis (Cain) S.I. Ahmed & Cain Aude: 22E/04 Sporormiella ovina (Desm.) S.I. Ahmed & Cain Aude: 17/02, 46/07 Sporormiella pascua (Niessl) S.I. Ahmed & Cain Aude/Pyrénées-Orientales: 20E/04 Sporormiella pulchella (E.C. Hansen) S.I. Ahmed & Cain Aude: 17E/02 Sporormiella teretispora S.I. Ahmed & Cain ex J.C. Krug Aude: 57/03; Aude/Pyrénées-Orientales: 22E/04; Aveyron: 111/97; Corsica: 6/01 Trichodelitschia bisporula (P. Crouan &H. Crouan) Munk Aude: 18E/04, 22E/04,1/05; Aude/Pyrénées-Orientales: 19E/04; Aveyron: 113/97; Cantal: 50/98; Corsica: 6/01, 8/01,Herault: 107/97, 108E/97, 115/97, 116/97, 26/98, 54/98, 55/98, 9/03, 10/03; Tarn: 20/98 Trichodelitschia munkii N. Lundq. Pyrénées-Orientales: 45/07

BASIDIOMYCOTINA Coprinellus curtus (Kalchbr.) Vilgalys,Hopple & Jacq. Johnson Ariège: 86/01 Coprinellus heptemerus (M. Lange &A.H. Sm.) Vilgalys,Hopple & Jacq. Johnson Aude: 30/05; Corsica: 5/01; Herault: 24/98, 52/98; Lot: 13/99; Tarn: 18/98 Coprinellus cf. heterosetulosus (Locq. ex Watling) Vilgalys,Hopple & Jacq. Johnson Aude: 46/07 Coprinellus marculentus (Britzelm.) Redhead,Vilgalys &Moncalvo Guadeloupe: 2E/07 Coprinellus pellucidus (P. Karst.) Redhead,Vilgalys &Moncalvo Aude/Pyrénées-Orientales: 20/04; Guadeloupe: 3E/07; Herault: 114/97, 115/97 Coprinopsis filamentifer (Kühner) Redhead,Vilgalys &Moncalvo Aude: 18/02, 12/03, 56E/03; Aveyron: 112/97; Herault: 110/97, 114/97, 115/97, 24/98 Coprinopsis nivea (Pers.) Redhead,Vilgalys &Moncalvo Guadeloupe: 4E/07, 5/07 Coprinopsis radiata (Bolton) Redhead,Vilgalys &Moncalvo Lot: 23/99; Morbihan: 15/99, Coprinopsis stercorea (Fr.) Redhead,Vilgalys &Moncalvo Aude: 5/02, 56/03, 46/07; Aude/Pyrénées-Orientales: 20/04; Corsica: 5E/01, 8/01; Guadeloupe: 1E/07; Herault: 109/97, 110/97, 114/97, 54/98, 8/03, 10/03; Lot-et- Garonne: 44/07; Tarn: 22/98, French coprophilous fungi 175

Coprinopsis utrifer (Joss. ex Watling) Redhead,Vilgalys &Moncalvo Aude: 11/03 Coprinopsis vermiculifer (Joss. ex Dennis) Redhead,Vilgalys &Moncalvo Aude: 18/02, 29/05, 30/05, 31/05 Coprinus cordisporus T. Gibbs Aude: 11E/03, 12/03, 57/03, 29/05; Aude/Pyrénées-Orientales: 20/04; Guadeloupe: 2E/07; Herault: 54/98 Coprinus ephemeroides (DC.) Fr. Aude: 12/03; Corsica: 9/01; Herault: 8/03 Coprinus cf. foetidellus P.D. Orton Aude: 46/07 Coprinus pseudoradiatus Kühner &Joss. Guadeloupe: 3E/07; Lot: 13/99 Panaeolus fimicola (Pers.) Gillet Aude: 29E/05 Panaeolus sphinctrinus (Fr.) Quél. Aude: 11E/03; Aude/Pyrénées-Orientales: 20E/04 Parasola misera (P. Karst.) Redhead,Vilgalys &Hopple Aude: 11/03, 12/03, 56E/03, 28/05, 46/07; Corsica: 6/01, 8/01, 9/01; Herault: 109/97, 115/97, 117/97, 24/98, 25/98, 26/98, 52/98, 56/98,8/03, 9/03, 10/03; Kerguelen: 4/01; Pyrénées-Orientales: 45/07 Together with Coprinopsis stercorea, one of the commonest and most widespread basidiomycetes occurring on incubated dung. Pholiotina coprophila (Kühner) Singer Aude: 29E/05 Psathyrella coprophila Watling Aude: 29/05 Psilocybe coprophila (Bull. ex Fr.) Quél. Aude: 19/02 Stropharia semiglobata (Batsch. ex Fr.) Quél. Herault: 51/98; Tarn: 19/98 MITOSPORIC FUNGI Dictyosporium toruloides (Corda) Guég. Herault: 114/97 Volutella ciliata (Alb. & Schwein.) Fr. Aude: 19/02; Lot-et-Garonne: 44/07

MYXOMYCOTA Didymium difforme (Pers.) Gray Lot-et-Garonne: 44/07

DISCUSSION

The sixty-seven samples from mainland France provided a total of 738 records of 150 species. The mean species richness of 11 per sample is within the range of values of 9-12 obtained for various herbivore mammalian dungs in the worldwide survey (Richardson 2001a). The five Corsican samples provided 176 M.J. Richardson

160 y = 9.32x0.652 140 R2 = 0.989

120 otal 100

80

60

40 Cumulative specie st 20

0 0 10 20 30 40 50 60 70 Sample number

Fig. 3. Cumulative total of taxa observed in successive samples of mammalian herbivore dung from mainland France and Corsica (latitude 41.75 – 44.75°N). The equation for the line of best fit is given on the graph.

a further 62 records of 41 species, of which seven were additional to those recorded from the mainland. The cumulative species curve for the 70 samples collected in a 3° belt of latitude in mainland France and Corsica (41.75-44.75°N) provide an estimate of 119 species to be expected from 50 samples (Fig. 3). Richardson (2001a, 2006) showed that studies of coprophilous fungi can be used to illustrate the latitudinal gradient of increasing species richness with decreasing latitude. The value of 119 species for around 43°N in France can be compared with values of 73-102 for samples from higher latitudes (65-50°N, Iceland, Canada/USA, Finland, Faroe Islands and the UK, and the Falkland Islands, 51oS), and 117-142 for lower latitudes (40°N-40°S, USA, Brazil, the Caribbean region,St Helena and Australia)(Richardson, 2006). The samples from the Kerguelen Islands produced a very limited mycota, with only 26 records of 13 species from nine samples. More samples would be needed to determine whether this is due to their remote position per se, or the difficulties of collecting and returning samples in a way that is not deleterious to the fungi. In contrast, the five samples from Guadeloupe provided 21 species typical of lower latitude coprophilous fungi, although the overall diversity estimate of 85 species from 50 samples from 21 Caribbean island samples was much lower than would be expected from tropical regions. It may be that this low value is due to the fact that the Caribbean islands have never been connected to a continental landmass, and have a restricted terrestrial mammalian fauna (Richardson, in press).

Acknowledgements. I am grateful to Jean-Louis Chapuis for obtaining samples from the Kerguelen Islands (as part of Programme no. 276 of the French Polar Institute), Andrew Richardson and Margo Thaller for collecting samples from Corsica and Ariège, and to Brian Cave for the samples from Morbihan and Lot. French coprophilous fungi 177

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